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Systematics of Simsia (Compositae-Heliantheae) Author(S): David M

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Systematics of Simsia (Compositae-Heliantheae) Author(S): David M Systematics of Simsia (Compositae-Heliantheae) Author(s): David M. Spooner Source: Systematic Botany Monographs, Vol. 30, Systematics of Simsia (Compositae- Heliantheae) (Nov. 12, 1990), pp. 1-90 Published by: American Society of Plant Taxonomists Stable URL: http://www.jstor.org/stable/25027790 Accessed: 08/07/2010 11:04 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=aspt. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Society of Plant Taxonomists is collaborating with JSTOR to digitize, preserve and extend access to Systematic Botany Monographs. http://www.jstor.org SYSTEMATICS OF SIMSIA (COMPOSITAE-HELIANTHEAE) David M. Spooner Vegetable Crops Research Unit Agricultural Research Service, U.S.D.A. Department of Horticulture University ofWisconsin Madison, Wisconsin 53706 ABSTRACT.Simsia (Compositae-Heliantheae) comprises annuals, herbaceous perennials, suffru tescent perennials, and shrubs. It occurs throughout tropical America from sea level to 3000 m. This study is based on comparative morphology, including extensive field studies and observations in the greenhouse, phenetic studies, crossing experiments, and cytology. Eighteen species including seven varieties are recognized. Two species, S. santarosensis and S. villasenorii, are newly described. The following new combinations are proposed: S. annectens var. grayi, S. foetida var. grandiflora, S. foetida var. jamaicensis, S. foetida var. megacephala, and S. foetida var. panamensis. Twenty species and varieties are predominantly self-incompatible and outcrossers; three species are self-compatible (or apomictic) and possibly inbreeders.Meiosis is normal inmost of the F1 hybrids produced from outcross ing species but irregular in F1 hybrids involving a self-compatible species. Chromosome numbers are uniformly n = 17. Counts are reported for the first time for the following: S. annectens var. annectens, S. chaseae, S. foetida var. panamensis, S. holwayi, S. molinae, S. santarosensis, S. setosa, S. tenuis, and S. villasenorii. INTRODUCTION Simsia is a genus of herbs and shrubs distributed from the southwestern United States to the center of Argentina. The greatest diversity occurs from central Mexico to Panama, where 18 of the 23 species and varieties grow. Some of the species (e.g., S. amplexicaulis, S. lagascaeformis) arewidespread roadside and agriculturalweeds and are familiar plants to collectors of Mexican Compositae. Others (e.g., S. holwayi, S. steyermarkii) are narrowly restricted endemics and are seldom encoun tered. Many of the taxa occur in dry, tropical deciduous forests (e.g., S. foetida), others in upland pine-oak to cloud forests (e.g., S. amplexicaulis); S. eurylepis occurs along the humid Mexican Gulf coastal lowlands. The species share a 2-4 seriate involucre, linear, sterile ray florets (except for one eradiate species, S. eurylepis), a convex receptacle with conduplicate, medium-textured, persistent pales, fertile disc florets with a well-defined, glandular-puberulent tube and abruptly expanded throat, and laterally flattened, ovate to obovate to elliptic disc achenes with or without awns (rarely with intermediate squamellae). The genus was last monographed by Blake (1913) who resurrected it from synonymy within Encelia Adanson and recognized 22 species and three varieties. In a recent useful and insightful synopsis of Mexican and Central American species, Robinson and Brettell (1972) treated 24 species from these areas, including seven new ones, and estimated 35 species total for the genus. This synopsis was based on limited material (only US), did not include field-investigations, and did not treat species from Jamaica and South America. Despite previous work on Simsia, basic taxonomic problems remain, as also 1 2 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 30 indicated by Nash (1976) and by Jansen and Stuessy (1980). The genus is taxonomi cally difficult by any standard. The polymorphic nature of many species has led to a confusing array of names and taxa of uncertain circumscription.A possible contrib uting factor to this taxonomic confusion is the apparent tendency of certain species to hybridize, a pattern discerned from herbarium investigations and further sug gested by field- and greenhouse-studies. The present study of Simsia is based on considerably more data than available to previous workers. Over 3600 herbarium specimens have been examined from 63 institutions. In addition, Viguiera Kunth and related genera have been examined at ENCB, GH, MEXU, MO, OS, and US for a study of generic relationships. Five field tripswere conducted over a total of eight months from 1982-1990 throughout Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Ecua dor, and Argentina; these resulted in field-observations and collections of all taxa except for one endemic variety from Jamaica. The field-investigations were espe cially useful for observations on intra- and interpopulational variability, habitats, natural hybridization, and for the collection of pertinent morphological observa tions difficult to obtain from herbarium collections, such as habit, plant height, branching patterns, corolla color, and leaf odor. Cytological collections provided material for 95 chromosome counts covering all taxa except S. foetida var. jamaicensis. Prior to this study, 56 chromosome counts were documented for 14 of the 23 species and varieties and one natural hybrid of Simsia; 93 new counts for 22 of these taxa and two natural hybrids (Appendix 1) are listed here. First counts are reported for S. annectens var. annectens, S. chaseae, S. foetida var. panamensis, S. holwayi, S. molinae, S. santarosensis, S. setosa, S. tenuis, S. villasenorii, and the natural hybrids S. am plexicaulis x S. lagascaeformis and S. foetida var. foetida x S. lagascaeformis. It is evident that Simsia has evolved at the diploid or diploidized level, and therefore speciation within the genus must have involved factors other than changes in ploidy level. Numerical phenetic studies have been used to help define taxonomic limits and to develop hypotheses on evolutionary patterns in the genus. Sixteen species and three additional varieties have been grown in the greenhouse; of these, 11 species and three varieties were used in an artificial crossing program. These crossing studies provided data on compatibility systems, genomic compatibility among vari ous species, stability of morphological features, and insights into the nature of taxonomic problems and evolutionary processes operative in Simsia. Simsia is here treated as comprising 18 species, including seven varieties. Two species, S. santarosensis and S. villasenorii, are newly described. The species are all monobasic (x = 17) and homoploid (2n = 34). Three species are self-compatible (or apomictic); all others are self-incompatible. Local hybridization is believed to occur between certain species. One species, S. chaseae, exhibits additive morphological features between S. foetida and S. eurylepsis that could have resulted either by hybrid speciation or by convergent evolution. TAXONOMIC HISTORY The taxonomic history of Simsia begins with the publication of Coreopsis foetida Cavanilles (1791), described from garden-grown plants raised from achenes 1990 SIMSIA 3 supplied by the Sesse andMocifio expedition. Persoon (1807) circumscribedSimsia, including S. foetida (Cavanilles) Persoon, and S. heterophylla (Cavanilles) Persoon [=Iostephane heterophylla (Cavanilles) Bentham ex Hemsley]. Kunth (1818) and Sprengel (1826) submerged Simsia intoXimenesia Cavanilles (= Verbesina L.), but Cassini (1829) recognized Simsia and greatly improved the characterization of the genus. Simsia was accepted as a genus by later workers, and new taxa were described by de Candolle (1836), Gray (1850, 1853), Bentham (1853), and Triana (1858). Gray (1873) later reduced Simsia to a section of Encelia. Subsequently, taxa now assigned to Simsia were described inEncelia (Hemsley 1881;Gray 1887;Rose 1895; Fernald 1897; Kuntze 1898;Hieronymus 1901;Greenman 1903;Millspaugh 1904). A modern interpretation of Encelia and related genera (or genera at least superfi cially similar to Encelia) awaited the insightfulwork of Blake (1913), who resur rected Simsia and described eight new taxa. Simsia has been accepted since that time, with later additions provided by Blake (1917; 1918a; 1924; 1928), Loesener (1923), Cuatrecasas (1954), and Robinson and Brettell (1972). GENERIC RELATIONSHIPS Simsia is relatively homogeneous, and, except for some close relatives in Viguiera, it rarely is
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